Search

Insect populations are affected by density-dependent and density-independent factors, and knowing how these factors affect long-term population growth is critical to pest management. In this study, we experimentally manipulated densities of the hemlock woolly adelgid on eastern and western hemlock trees in the western USA to evaluate the effects of density and host species on hemlock woolly adelgid crawler colonization. We then followed development of hemlock woolly adelgid on each hemlock species. Settlement of crawlers was strongly density-dependent and consistent between host species. In addition, a period of hot days that coincided with the settlement of hemlock woolly adelgid crawlers put our experimental and naturally occurring populations into diapause during April. Diapause resulted in one generation that yr in our experimental population. Analyses of long-term air temperature records indicated that diapause-inducing temperatures in April similar to those observed in our experiment have occurred rarely since 1909 and the frequency of these events has not changed over time. Prior work suggests that hemlock woolly adelgid completes two generations per yr in the western USA with a diapause occurring in the summer. This typical life history reflects the long-term influence of regional average seasonal temperature patterns on development and the timing of diapause-inducing temperatures. However, the timing of unseasonal weather, such as the hot days observed in our experiment, occasionally changes life history trajectories from this normal pattern. Our results show that density-dependent and density-independent factors have strong effects on generational mortality and life history of hemlock woolly adelgid that are important to its population dynamics and management., (Published by Oxford University Press on behalf of Entomological Society of America 2016. This work is written by US Government employees and is in the public domain in the US.)

Varying densities of the spring generation of the hemlock woolly adelgid were manipulated on 16 previously uninfested eastern hemlocks in an open-field plantation in Massachusetts. In contrast to experimentally created hemlock woolly adelgid populations in a forest, as reported previously, there was no evidence of density-dependent survival on a tree-wide basis in the plantation in the spring (progrediens) generation. There was, however, comparable density-dependent survival of settled crawlers and sexupara production when samples of the population were examined from branches with high density. Plantation hemlocks had 9.3 times more foliage and 10 times lower adelgid densities per cm than the forest hemlocks. These results show that density-dependent processes in the progrediens generation may only be evident when hemlock woolly adelgid density reaches a critical threshold. In the sistens generation that begins in midsummer, we counted a mean of 126 settled crawlers on marked branch on all 16 trees, but none of these adelgids survived the mid-summer aestivation phase, perhaps due to higher temperatures that were recorded in the plantation compared with a nearby hemlock forest, where 16% of the adelgids survived the aestivation phase. Whole tree counts of overwintering adelgids revealed that the adelgid populations had gone extinct on 13 out of the 16 trees. Mortality in the midsummer aestivation phase often exceeds overwintering mortality that has been widely thought to be the main factor that limits adelgid population growth and spread, particularly in northern states., (© The Author 2015. Published by Oxford University Press on behalf of Entomological Society of America. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

The hemlock woolly adelgid (Adelges tsugae Annand) is an invasive species causing high mortality of eastern hemlock (Tsuga canadensis (L) Carriere) in the forests of eastern North America. Recent findings revealed that sapling eastern hemlocks artificially infested with hemlock woolly adelgid in a plantation setting responded to the insect with an array of induced resin volatile changes. Here we determine if eastern hemlocks growing beneath a forest canopy respond to hemlock woolly adelgid infestation with the same patterns of constitutive and inducible volatile resin production as those plantation specimens. We inoculated previously uninfested branches of mature and immature hemlocks in a central New England forest with hemlock woolly adelgid. We then sampled twig tissue of infested and uninfested trees in late spring, early summer, and mid-autumn, after known intervals of adelgid activity when an induced response might be expected. We identified and quantified resin volatiles by gas chromatography-mass spectrometry. Constitutive levels of total monoterpenoids, sesquiterpenoids, and combined resin volatiles were all several-fold more abundant in forest trees than those previously measured in a plantation setting, pointing to further study of the influence of site factors on hemlock volatile production. Hemlock woolly adelgid infestation induced an array of changes in eastern hemlock's volatile profile, including many-fold increases in benzyl alcohol and methyl salicylate accumulation. Despite differences in constitutive concentrations of volatiles between the two sites, our findings verify that hemlock woolly adelgid elicits patterns of resin volatile induction in forest-grown eastern hemlocks quite similar to those previously observed in plantation grown trees.

The hemlock woolly adelgid (Adelges tsugae Annand) has decimated eastern hemlocks (Tsuga canadensis Carrière) in forests throughout the eastern United Sates, but its densities in central New England appear to have stabilized. To find out why, we infested 64 eastern hemlocks with varying densities of adelgid ovisacs in a typical eastern hemlock forest in western Massachusetts. We subsequently documented adelgid density, fecundity, and the amount of new growth on experimental trees over two consecutive years. We used a 2 by 2 randomized block design using previously and newly infested hemlocks that were either 1-m tall saplings or branches of mature trees. There was a density-dependent decline in the survival and fecundity of adelgid in both the spring and winter generations. This response was a function of both previous infestation by adelgid and current year's crawler density in the spring generation. Additionally, the production of sexuparae in the spring generation played a key role in the overall density-dependent survival of adelgid, suggesting that sexuparae production is strongly linked to developing crawler density.

Recent studies have identified a small number of individual eastern hemlock trees that demonstrate relative resistance to the introduced sap-feeding insect, the hemlock woolly adelgid. Using gas chromatography, we compared concentrations of twenty-two terpenoids in susceptible and relatively resistant trees, both in the forest and in propagated cuttings in a common-garden setting. Terpenoid concentrations were higher in twig tissue of resistant versus susceptible trees, across six sampling dates and at both sites. These changes may be particularly important because the hemlock woolly adelgid feeds on twig tissue. Because the common-garden cuttings were free of herbivores, the higher terpenoid concentrations are interpreted as a constitutive defense. Increased levels of monoterpenes and sesquiterpenes imply an overall increase in the input of carbon precursors to both terpenoid synthesis pathways.

Monitoring programs increasingly are used to document the spread of invasive species in the hope of detecting and eradicating low-density infestations before they become established. However, interobserver variation in the detection and correct identification of low-density populations of invasive species remains largely unexplored. In this study, we compare the abilities of volunteer and experienced individuals to detect low-density populations of an actively spreading invasive species, and we explore how interobserver variation can bias estimates of the proportion of sites infested derived from occupancy models that allow for both false negative and false positive (misclassification) errors. We found that experienced individuals detected small infestations at sites where volunteers failed to find infestations. However, occupancy models erroneously suggested that experienced observers had a higher probability of falsely detecting the species as present than did volunteers. This unexpected finding is an artifact of the modeling framework and results from a failure of volunteers to detect low-density infestations rather than from false positive errors by experienced observers. Our findings reveal a potential issue with site occupancy models that can arise when volunteer and experienced observers are used together in surveys.

Although biological invasions are of considerable concern to ecologists, relatively little attention has been paid to the potential for and consequences of indirect interactions between invasive species. Such interactions are generally thought to enhance invasives' spread and impact (i.e., the "invasional meltdown" hypothesis); however, exotic species might also act indirectly to slow the spread or blunt the impact of other invasives. On the east coast of the United States, the invasive hemlock woolly adelgid (Adelges tsugae, HWA) and elongate hemlock scale (Fiorinia externa, EHS) both feed on eastern hemlock (Tsuga canadensis). Of the two insects, HWA is considered far more damaging and disproportionately responsible for hemlock mortality. We describe research assessing the interaction between HWA and EHS, and the consequences of this interaction for eastern hemlock. We conducted an experiment in which uninfested hemlock branches were experimentally infested with herbivores in a 2 x 2 factorial design (either, both, or neither herbivore species). Over the 2.5-year course of the experiment, each herbivore's density was approximately 30% lower in mixed- vs. single-species treatments. Intriguingly, however, interspecific competition weakened rather than enhanced plant damage: growth was lower in the HWA-only treatment than in the HWA + EHS, EHS-only, or control treatments. Our results suggest that, for HWA-infested hemlocks, the benefit of co-occurring EHS infestations (reduced HWA density) may outweigh the cost (increased resource depletion).